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<I>Callinectes Ornatus</I> (Brachyura, Portunidae)

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<I>Callinectes Ornatus</I> (Brachyura, Portunidae) BULLETIN OF MARINE SCIENCE, 46(2): 274-286,1990 MORPHOMETRY AND SIZE AT MATURITY OF CALLINECTES ORNATUS (BRACHYURA, PORTUNIDAE) IN BERMUDA Paul A. Haefner, Jr. ABSTRACT Morphometry and size at maturity are described for 196 specimens of Ca/linectes ornatus collected in Mullet Bay, Bermuda during summers of 1981 and 1987. Carapace width in- cluding and excluding lateral spines, body depth, and male abdomen width exhibited linear and isometric growth relative to short carapace length (CSL), the independent variable. Female abdomen width and chelar propodallength, width, and depth were allometric. Prepubertal (near 20 mm CSL) and pubertal molts (near 30 mm CSL) of males were inferred from changes in allometry of chelar propodal dimensions. The prepubertal molt of females (near 24 mm CSL) was revealed by an allometric change in abdominal width and cheliped dimensions. Allometric changes were correlated with morphological changes in the abdomen and pleopods and with gonadal development. Females attain puberty (near 45 mm CSL) at terminal ecdysis as crabs metamorphose to the adult configuration. The species is heterochelic and hetero- dontic. The molariform claw occurred on the right side of 83.5% of all crabs (no significant difference between sexes). Cheliped laterality changes with age; frequency of "right-hand- edness" decreases with increasing size (age). Highly significant differences occurred between molariform (right) and serratiform (left) chelipeds for all dimensions for both sexes. Few papers have been published on the biology of the Brachyura of Bermuda since Verrill's (1908) monograph. Markham and McDermott (1980) listed 106 Bermudan brachyurans, and Chace et al. (1986) provided diagnostic and distri- butional comments for 55 of those species, but the natural history and reproductive biology of many of those species have not been described. Callinectes ornatus is a relatively large portunid crab closely related to the commercial blue crab, C. sapidus, of the Atlantic and Gulf coasts of North America (Williams, 1974). Although C. ornatus might be of potential commercial value in Bermuda, its significance is more likely dependent upon its ecological role as scavenger, predator, and food resource for other aquatic organisms and shore birds. C. ornatus ranges from North Carolina to Brazil, including the Caribbean is- lands, and Bermuda (Verrill, 1908). Although records indicate tolerance to a broad salinity range, most collections have been made in warm (18-3 1°C), high salinity (18-350/00)water (Williams, 1974; Norse, 1978). Spawning probably occurs year- round, based on evidence from museum specimens of ovigerous females collected in nearly every month (Williams, 1974). Seasonal abundances of C. ornatus on seagrass beds in the Indian River lagoon in Florida (Gore, 1977) and of C. arcuatus and C. toxotes on the Mexican Pacific coast (Paul, 1982) appear to be related to seasonal changes in rainfall; in each instance, maximum abundance occurred during the dry season. Paul (1982) observed distinct spawning seasons and some segregation of the sexes within the estuarine lagoon system for both species. This paper describes the morphometry and size at maturity of C. ornatus in Mullet Bay, Bermuda. METHODS All specimens were collected in Mullet Bay, a Thalassia embayment opening into Ferry Reach in St. Georges' Parish, Bermuda. A long-handled, "D-shaped" dipnet was used to collect crabs in the 274 HAEFNER: MORPHOMETRY OF CALLINECTES ORNATUS 275 lower intertidal zone within one hour of low tide. Collections in 1981 were made during the first three weeks of August; in 1987, they were made from 17 June to 17 July. All specimens were returned to the laboratory and either refrigerated or maintained in running seawater prior to examination. The following measurements were made on each specimen: carapace width including (CLW) and excluding lateral spines (CSW); carapace length excluding frontal spine (CSL); body depth (BD); abdomen width of the fourth and sixth segments (FAW4, FAW6) in females, and the fused third-fifth segment of males (MAW); propodal dimensions of the right and left chelipeds: length (RCPL, LCPL),. width (RCPW, LCPW), and depth (RCPD, LCPD). Fresh, entire, intact crabs were blotted dry and weighed (WT) to the nearest 0.1 g. The abdomens and pleopods were examined to assess maturity. I recognized four progressive stages of development of the male abdomen. In the immature stage, the abdomen is sealed in a flexed position to the sternum, and the pleopods collectively are enclosed within a transparent tissue sheath. A prepubertal abdomen is sealed, but the pleopods are unsheathed. In the pubertal stage, the abdomen is unsealed but partially locked to the sternum by paired tubercles, similar to the device of C. sapidus (Pyle and Cronin, 1950). The penis is inserted in the gonopods. An adult male has a free, fully extensible abdomen. A female abdomen is considered immature if it is sealed and the vulvae are indistinct. In the prepubertal stage, the vulvae are evident, but the abdomen is sealed to the sternum. The penultimate instar female possesses a triangular, lightly pigmented abdomen. The adult female has the characteristic broadened, heavily pigmented and fully extensible abdomen, and setose pleopods. Squash preparations of vasa deferentia were observed microscopically for existence of spermato- ph ores, a sign of sexual maturity (Haley, 1973). Squash preparations of ovaries were examined mi- croscopically to determine stages of development (Pradeille-Rouquette, 1976; 1978). Samples of fluid were withdrawn from incisions in the female spermathecae and examined microscopically for presence of sperm or spermatophores. Plots of each measurement relative to CSL, the selected reference variable, were examined for linearity. Curvilinear relationships were subjected to bivariate analysis in which bivariate ratios (Y/CSL) were determined for each of the suspect variables. Discontinuities in the series reveal changes in the rate of growth ofa given dimension and often are indicative of size ranges within which crabs undergo pubertal and maturational molts (Gore and Scotto, 1983; Davidson and Marsden, 1987). Crabs were grouped into 5 mm (CSL) size classes, and cross tabulations (contingency tables) of developmental stages of abdomen, pleopods, and gonad were tested by Chi-square analysis. These data arrays were compared with the bivariate discontinuities to assess the reliability of the bivariate analyses in predicting stages of sexual maturity. Linear least squares regression equations were derived for each parameter for the total sample of crabs as well as for data subsets which were selected on the basis of gonadal and bivariate analyses. Allometry of growth was examined by applying the equation CSL = aXb to logarithmically transformed data, where (a) is the intercept, and b is the allometric constant. Randomness ofheterodentition and heterochelation with respect to handedness was tested by Chi- square analysis. The dimensions of the major (molariform) chelae were compared with those of the minor (serratiform) chelae by matched pair analysis. RESULTS The relationships between CSL and the dimensions CLWand CSW are linear and isometric (Table 1).The BD:CSL relationship is linear with positive allometry. Sexual dimorphism is not evident in these dimensions. Covariance analysis of restricted data sets (inclusion of males and females within the same CSL range) revealed heterogeneity of variance for two of the regressions tested (Table 2). Although this normally precludes use of the pooled equation to describe the total population for the dimension in question (Snedecor and Cochran, 1967), the marginally significant test of variance, coupled with non-significance of the tests for regression and intercept, support use of the pooled equations for CLW, CSW and BD. The log-log equation for the weight-length relationship (Table 1)is an acceptable model and, for reasons stated above, the pooled equation is valid even though heterogeneity of variance was revealed by covariance analysis (Table 2). The male abdomen width:CSL relationship is linear and isometric (Table 1). The female abdomen width:CSL relationships differ from each other (Table 1). 276 BULLETIN OF MARINE SCIENCE, VOL. 46, NO.2, 1990 Table I. Regression analyses of body dimensions of Ca/linecles ornatus. based on short carapace length (CSL) as independent variable. Adult female excluded from analyses. Constant of allometry (b), based on log-log transformations, provided for all relationships. Asterisks indicate positive allom- etry Y variable Sex N Least square regression r' x 100 b CLW M 116 y= 1.078 + 2.096X 99.58 0.988 CLW F 75 Y = -0.209 + 2.159X 99.35 1.013 CLW M+F 190 y= 0.880 + 2.109X 99.50 0.993 CSW M 114 Y = -0.926 + 1.742X 99.62 1.034 CSW F 74 Y = -0.819 + 1.734X 99.72 1.026 CSW M+F 186 Y = -0.912 + 1.740X 99.65 1.031 BD M liS Y = -0.511 + 0.590X 99.73 1.048* BD F 76 Y = -0.453 + 0.586X 99.65 1.060* BD M +F 189 Y = -0.516 + 0.590X 99.73 1.051 * WT M 105 LOG Y = -3.415 + 3.163 LOG X 99.83 WT F 57 LOG Y = -3.379 + 3.133 LOG X 99.79 WT M +F 160 LOG Y = -3.411 + 3.159 LOG X 99.81 MAW M 118 y= 0.694 + 0.496X 99.47 0.960 FAW4 F 73 Y = -0.868 + 0.607X 98.27 1.079* FAW6 F<24mmCSL 51 Y= -1.416 + 0.319X 96.32 1.409* FAW6 24-38 mm CSL 22 Y = -3.676 + 0.414X 84.44 1.455* Although both are allometric, the relationship of FA W4:CSL is linear (Fig. 1) whereas FAW6:CSL is curvilinear (Fig. 2). Bivariate ratio analysis of the FAW6: CSL relationship revealed a discontinuity in the series, spanning the 21-27 mm CSL size range. For that reason the data were divided into two arrays with 24 mm CSL selected as the size of transition (Table 1).
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